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Worms and Human Disease Second Edition R Muller
Digital Instant Download
Author(s): R Muller
ISBN(s): 9780851995168, 0851995160
Edition: Second
File Details: PDF, 4.95 MB
Year: 2002
Language: english
00Worms & H.D. - Prelims. 20/11/01 1:41 PM Page i
and
Former Director
International Institute of Parasitology
St Albans, Hertfordshire, UK
CABI Publishing
00Worms & H.D. - Prelims. 14/11/01 4:24 PM Page iv
Contents
Acknowledgements ix
Introduction 1
1. The Trematodes 3
Morphology 5
Life Cycle Stages 6
Classification 6
Family Schistosomatidae 9
Schistosomes 9
Family Paragonimidae 32
Paragonimus westermani 32
Family Achillurbainiidae 37
Family Opisthorchidae 38
Clonorchis sinensis 38
Opisthorchis viverrini 43
Opisthorchis felineus 44
Family Dicrocoeliidae 45
Dicrocoelium dendriticum 45
Family Fasciolidae 46
Fasciola hepatica 46
Fasciolopsis buski 49
Family Heterophyidae 51
Heterophyes heterophyes 51
Metagonimus yokogawai 53
Family Paramphistomidae 55
Gastrodiscoides hominis 55
Family Echinostomidae 56
Echinostoma ilocanum 56
Other Echinostomids 56
Other Occasional and Rare Human-parasitic Trematodes 58
Family Diplostomidae 58
Family Lecithodendriidae 58
Family Plagiorchiidae 58
Family Troglotrematidae 58
Other families 59
v
00Worms & H.D. - Prelims. 14/11/01 4:24 PM Page vi
vi Contents
2. The Cestodes 63
Classification 64
Order Pseudophyllidea 65
Diphyllobothrium latum 65
Sparganosis 70
Order Cyclophyllidea 71
Family Taeniidae 71
Taenia saginata 71
Taenia solium 76
Cysticercosis 80
Taenia multiceps 83
Echinococcus granulosus 85
Echinococcus multilocularis 94
Echinococcus oligarthrus 97
Echinococcus vogeli 97
Family Hymenolepididae 98
Hymenolepis nana 98
Hymenolepis diminuta 101
Family Dipylididae 102
Dipylidium caninum 102
Very occasional human tapeworms 102
Contents vii
viii Contents
Index 287
00Worms & H.D. - Prelims. 14/11/01 4:24 PM Page ix
Acknowledgements
ix
00Worms & H.D. - Prelims. 14/11/01 4:24 PM Page x
01Worms Intro. & Chap 01 14/11/01 4:24 PM Page 1
Introduction
While this book is the second edition of concerned with tropical and exotic dis-
Worms and Disease: a Manual of Medical eases and for students taking postgraduate
Helminthology (1975), because of the long degrees and diplomas in aspects of tropical
time that has elapsed since the publication and infectious diseases. It should also
of the earlier book, it has been so exten- prove useful as an accessory text and refer-
sively revised and brought up to date that ence source for undergraduate medical,
virtually every chapter has had to be zoological and tropical health engineering
almost completely rewritten. In the inter- students, and for medical technologists,
vening years the importance to humans of microbiologists and physicians in temper-
some new helminths has emerged, such as ate climates. With increase in air travel,
Oesophagostomum bifurcum and Para- most hospitals and medical practitioners in
strongylus costaricensis, but principally developed countries are meeting cases of
the changes have been necessitated by the parasitic infections that may have been
great strides that have been made in knowl- very rare occurrences in the past, and it is
edge of the diagnosis, treatment, immunol- becoming increasingly necessary to ask of
ogy and molecular biology of parasites. The almost all patients ‘Unde venis?’. Also, if
chapter on the immunology of helminths global warming increases, it is likely that
(now written by Derek Wakelin) has been the endemicity of some helminth infec-
greatly amplified, with the addition of tions will extend to higher latitudes.
more detailed paragraphs in the appropri- The format of this book is fairly conven-
ate sections, together with the latest infor- tional, with parasites considered in order
mation on the prospects for specific of their zoological relationships rather than
vaccines. There has also been exciting their location in the body. The latter
progress in the field of global control of approach may be useful for diagnosis but is
various helminths, such as the schisto- not practical for other aspects of the sub-
somes, soil-transmitted nematodes, filariae ject, as some parasites can occupy a wide
(both those causing lymphatic filariasis and range of sites in the body, so that there
those causing onchocerciasis) and the would be a great deal of repetition, and
guinea worm. Most of these campaigns also because the relationships between
have been possible because of recent many similar parasites that occupy differ-
advances in chemotherapy and, in some ent organs would be obscured. However,
cases, of diagnosis; many have been linked the various possible locations in the body
with efforts to improve sanitation and of all the important helminths are shown
general health. in Fig. 132 and alternative diagnoses are
The book is intended principally as a discussed in the appropriate individual
practical guide in human helminthology sections. An attempt has also been made to
for physicians and medical technologists have the best of both worlds, e.g. all the
1
01Worms Intro. & Chap 01 14/11/01 4:24 PM Page 2
2 Introduction
Adult trematodes, or flukes, may be found means synonymous with the presence of
in the intestinal tract, bile-ducts, lungs or disease. In contrast to viruses, bacteria or
blood of humans. Some details concerning protozoans, trematodes do not multiply
the medically most important species are within the human body and the few organ-
shown in Table 1. All the trematodes men- isms present in the great majority of
tioned in the table are normal human para- infected persons are tolerated with the
sites, except some species of Paragonimus minimum of inconvenience and are often
and Fasciola and some heterophyids and not diagnosed. It is the small percentage of
echinostomes, which are accidental para- patients with large worm burdens (so-
sites with humans not being involved in called ‘wormy people’) or in whom the par-
their transmission cycles. However, almost asites or their eggs are in ectopic sites in
all trematodes are very catholic in their the body who give cause for alarm.
choice of definitive hosts (a notable excep- The digenetic trematodes are members
tion is Schistosoma haematobium) and of the phylum Platyhelminthes, which also
have a wide range of animal reservoirs; 144 includes the cestodes (tapeworms), mono-
species that have been found in humans geneans (ectoparasites of fishes and
are mentioned in the text, most of which amphibians) and free-living turbellarians
are natural animal parasites. Not shown in (planarians, etc.). Platyhelminthes, or flat-
the table are various aberrant forms, such worms, are acoelomate bilateria (bilaterally
as the cercarial larvae of animal and bird symmetrical and lacking a coelom). The
schistosomes, which can penetrate the skin excretory system is based on the flame cell,
of humans but are not able to mature. or protonephridium, and often the pattern
Pre-eminent in medical and economic of flame cells can be of importance in clas-
importance are the schistosomes, or blood sification. Trematodes are characteristically
flukes, which are the cause of one of the flat and leaflike, or occasionally globular,
major human diseases, schistosomiasis. hermaphroditic organisms (except for the
This is a source of suffering in many warm schistosomes, which have a male folded
countries and is a major cause of morbid- about its long axis and a cylindrical female
ity. No other trematode is the cause of such (Figs 3 and 4)). All have complicated life
widespread morbidity, but liver flukes cycles with alternating sexual and asexual
(Clonorchis and the closely related development in different hosts. Asexual
Opisthorchis) and lung flukes (Paragonimus) multiplication takes place in a snail, and
are important parasites in areas of Asia and for parasites of medical importance this is
their presence may result in severe disease always a gastropod snail. It is believed that
and possibly death. the trematodes were originally parasites of
It needs to be stressed that the presence molluscs and they are still always very
of trematode parasites in the body is by no specific in their choice of snail host;
3
4
Table 1. Trematodes of medical importance.
Chapter 1
Situation of Eggs recovered Snail intermediate Other intermediate Geographical
Habitat Species adult from host or transport hosts distribution
Blood Schistosoma Mesenteric veins Faeces Biomphalaria spp. None (active penetration Africa, South America
mansoni by cercariae)
S. japonicum Mesenteric veins Faeces Oncomelania spp. None (active penetration China, South-East Asia
by cercariae)
S. mekongi Mesenteric veins Faeces Neotricula None (active penetration Cambodia, Laos
by cercariae)
S. intercalatum Mesenteric veins Faeces Bulinus spp. None (active penetration Central Africa
by cercariae)
S. haematobium Vesicular veins Urine Bulinus spp. None (active penetration Africa, Middle East
by cercariae)
Lungs Paragonimus Cysts in lungs Sputum and faeces Semisulcospirura Edible crustaceans South-East Asia,
westermani Thiara containing China, Japan
Oncomelania metacercariae
Paragonimus spp. Cysts in lungs Sputum and faeces Various Edible crustaceans South-East Asia,
containing West Africa, South and
metacercariae Central America
Liver Clonorchis sinensis Bile and pancreatic Faeces Bulimus Freshwater food fish South-East Asia
ducts Parafossarulus containing metacercariae
Opisthorchis felineus Bile and pancreatic Faeces Bithynia Freshwater food fish Siberia, East Europe
ducts containing metacercariae
O. viverrini Bile and pancreatic Faeces Bithynia Freshwater food fish Thailand, Laos
ducts containing metacercariae
Fasciola hepatica Bile ducts Faeces Lymnaea Metacercariae encysted Cosmopolitan (mainly
on plants temperate areas)
Intestine Fasciolopsis buski Small intestine Faeces Segmentina Metacercariae on South-East Asia, India
water plants
Heterophyes Small intestine Faeces Pirenella Freshwater food fish South-East Asia,
heterophyes Cerithidea containing metacercariae Middle East, Egypt,
southern Europe
Metagonimus Small intestine Faeces Semisulcospira Freshwater food fish South-East Asia, Russia
yokogawai containing metacercariae (Siberia), southern Europe
Other heterophyids Small intestine Faeces Various Freshwater food fish Worldwide in warm
containing metacercariae countries
Echinostomes Small intestine Faeces Various Freshwater fish or snails Mostly South-East
containing metacercariae Asia, India
Gastrodiscoides Caecum and colon Faeces Helicorbis Metacercariae on South-East Asia
hominis water plants
The Trematodes 5
Fig. 1. Snails that act as first intermediate hosts of the trematodes of medical importance.
(a) Biomphalaria glabrata from Brazil, host of Schistosoma mansoni. (b) Bulinus (Physopsis) globosus
from Nigeria and (c) Bulinus (Bulinus) truncatus from Iran, hosts of S. haematobium. (d) Oncomelania
hupensis nosophora from Japan, host of S. japonicum. (e) Thiara granifera from China and
(f) Semisulcospira libertina from China, hosts of Paragonimus westermani and Metagonimus yokogawai.
(g) Polypylis hemisphaerula from China, host of Fasciolopsis buski. (h) Parafossarulus manchouricus
from China, host of Clonorchis sinensis. (i) Codiella (= Bithynia) leachi from Germany, host of
Opisthorchis felineus. (j) Pirenella conica (from Egypt) host of Heterophyes heterophyes. (k) Lymnaea
trunculatula from England, host of Fasciola hepatica.
6 Chapter 1
Classification
future (Brooks et al., 1985; Rohde et al., The modes of infection of trematodes
1993), but changes are not generally of medical importance shown in Table 2
accepted. Only a very few of the numerous reflect quite well the taxonomic divisions
families comprising the subclass Digenea are (the odd one out being Echinostoma, in
included (Yamaguti, 1971) – those which which it might be expected that the
have members of medical importance. cercariae would encyst on vegetation).
Fig. 3. Diagrams of the shape and principal organ systems of the hermaphrodite trematodes of medical
importance (schistosomes are shown in Fig. 4). Comparative sizes shown in silhouette.
8 Chapter 1
Schistosoma
Fasciolopsis
Fasciola
Gastrodiscoides?
SUPERFAMILY DICROCOELIOIDEA
Cercariae encyst in arthropods and have an
oral stylet. Family Schistosomatidae
Family Dicrocoeliidae (MBRA)
Found in intestine, liver, gall-bladder and Schistosomes
pancreas. The oral sucker is subterminal.
Testes adjacent or in tandem, anterior to At least seven species are parasites of
ovary. Vitellaria posterior to ventral humans: Schistosoma haematobium
sucker. (Bilharz, 1852); Weinland, 1858; S. mansoni
Dicrocoelium Sambon, 1907; S. japonicum Katsurada,
1904; S. intercalatum Fischer, 1934; S.
SUPERFAMILY OPISTHORCHIOIDEA (MBR) malayensis Greer, Ow-Yang and Yong,
Cercariae encyst in or on fish. 1988; S. mekongi Voge, Bruckner and
Family Opisthorchiidae (MB) Bruce, 1978; S. sinensium Pao, 1959.
Suckers weak. Semi-transparent flukes
found in bile-ducts and gall-bladder. Testes
in tandem behind ovary.
SYNONYMS (for S. haematobium)
Clonorchis, Opisthorchis, Metorchis
Distoma haematobia Bilharz, 1852;
Family Heterophyidae (MB) Bilharzia haematobium Diesing, 1859.
This is a large family. All members are
potential parasites of humans. They are
minute flukes with a spinose tegument. LOCAL NAMES
Testes adjacent behind ovary. Au chung (Chinese), Tsagiya (Hausa, S.
Heterophyes, Metagonimus haematobium), Katayamabayo, Suisho-
SUPERFAMILY PLAGIORCHIOIDEA choman or Harapari (Japanese), Laremo
Family Lecinthodendriidae (MBRA) (Luo, S. haematobium haematuria), Kadi
Small spiny flukes with gonads in fore- dhig (Somali, S. haematobium haema-
body. Oral sucker large, ventral sucker turia), Kichocho (Swahili), Pa-yard bai-
small. Metacercariae in aquatic insects. mai lohit (Thai), Atosi eleje (Yoruba, S.
Phaneropsolus, Prosthodendrium haematobium).
10 Chapter 1
*This is the standardized nomenclature for parasitic diseases advocated by the World Association for
the Advancement of Veterinary Parasitology (WAAVP) (Kassai et al., 1988. Veterinary Parasitology 29,
299–326) and the World Federation of Parasitologists (WFP), but has not been widely adopted in
medical helminthology, particularly by the World Health Organization (WHO). For instance, of the titles
of references in this book, 112 have -iasis endings and 16 -osis, most of which are mentioning
cysticercosis or echinococcosis – used in both systems.
The Trematodes 11
S. japonicum (S. malayensis and S. ium. Eggs of each species can be recognized
mekongi). The male measures 12–20 mm by differences in size and morphology
0.5–0.55 mm and has no cuticular tubercu- (Table 3 and Fig. 6). On immersion in fresh
lations. There are 6–7 testes. The female water, particularly under conditions of
has the ovary at about the middle of the warmth and light, they hatch almost imme-
body. There are 5–200 eggs in the uterus at diately. The miracidium larvae (Fig. 7)
one time. swim actively by means of the cilia with
which they are covered and attempt to pen-
S. intercalatum. The male measures etrate any freshwater snail they come into
11–14 mm 0.3–0.4 mm. There are 2–7 contact with. The miracidia die in 16–32 h
testes. The female has 5–60 eggs in the if they do not succeed in reaching a suitable
uterus at one time. snail intermediate host. Like all trematodes
Of the complete 270 Mb genome of they are extremely host-specific in regard to
Schistosoma, 18–24% has so far been the snails in which they will develop, often
deposited in a database (Williams and far more so than in the definitive host. The
Johnston, 1999). species of snail parasitized depends on the
geographical region, but S. haematobium
LIFE CYCLE and S. intercalatum develop in snails of the
The eggs are passed in urine in S. haemato- genus Bulinus, S. mansoni in Biomphalaria
bium and in the faeces in all the other and S. japonicum in Oncomelania.
species and contain a fully formed miracid- Oncomelania differs from the other two
12 Chapter 1
Situation in human Mesenteric veins Mesenteric veins Vesical veins Mesenteric veins Mesenteric veins
Male
Length (mm) 10–20 6–12 10–14 11–14 15
Width (mm) 0.5 1.1 0.9 0.3–0.4 0.4
No. of testes 6–7 4–13 (usually 6–9) 4–5 2–7 (usually 4–5) 6–9
Tuberculations None Coarse Fine Fine None
Caecal junction Posterior third of body Anterior third of body Middle Middle Posterior third of body
Female
Length (mm) 20–30 10–20 16–20 10–14 12
Width (mm) 0.3 0.16 0.25 0.15–0.18 0.23
Uterus Anterior half of body Anterior half of body Anterior two-thirds of body Anterior two-thirds of body Anterior half of body
Number of eggs in
uterus 50–200 1–2 10–50 5–60 10+
Position of ovary Middle Anterior third of body Posterior third of body Posterior half of body Posterior half of body
Mature egg shape Round with small knob Lateral spined Terminal spined Terminal spined Round, small knob
and mean size 85 µm 60 µm 140 µm 61 µm 150 µm 62 µm 176 µm 61 µm 57 µm 66 µm
Mode of voiding eggs Faeces Faeces Urine Faeces Faeces
Egg production per 3500 100–300 20–300 150–400 ?
female per day (in
experimental animals)
Reaction of egg to Positive Positive Negative Positive Positive?
Ziehl–Neelsen stain
Intermediate Oncomelania Biomphalaria Bulinus Bulinus Tricula
host snail
The Trematodes
Distribution overlap None Over most of Africa Throughout range None
13
14 Chapter 1
Fig. 4. Diagram of the structures of the three major schistosome species. Only the male of
S. mansoni is shown. The males of the other species differ principally in the number of testes, while
that of S. japonicum has a smooth tegument.
Fig. 5. Integument and double outer membrane of S. mansoni. Electron micrograph. Original
magnification 83,500.
venous vessels and are carried to the lungs pairs. The schistosomula are usually
4–7 days after penetration (Figs 9 and 10). assumed to travel to the liver via the blood
The schistosomula move from the lungs to system (against the blood flow) but, at least
the portal vessels and there grow into adult for those of S. japonicum, some penetrate
schistosomes, which mate and remain in directly through the diaphragm. The adult
Fig. 6. Eggs of (a) S. haematobium, (b) S. mansoni and (c) S. japonicum. Actual size of (a) is 125 µm.
The Trematodes
15
16 Chapter 1
Fig. 7. Ciliated miracidium of S. mansoni. Note anterior penetration glands. Dark field. Actual size 80 m.
HUMAN
ADULTS mature
in 6–12 weeks.
In veins of bowel:
S.m., S.j., S.i.
In veins of bladder:
S.h.
liver
Some EGGS retained
lungs in tissues
lymph and veins
to heart
SCHISTOSOMULA
in skin
EGGS
Male and female In urine: S. haematobium
CERCARIAE In faeces: all others
penetrate skin
SNAIL S.m.
CERCARIAE emerge
Asexual Biomphalaria spp. .
reproductive S .j
SECONDARY
SPOROCYSTS stages in
i.
S.
gland weeks
an
h.
S.
MIRACIDIA
penetrate snail
Bulinus spp.
Fig. 9. The life cycle of schistosomes. Figure from Book of the Dead papyrus. S.m., S. mansoni; S.j.,
S. japonicum; S.h., S. haematobium; S.i., S. intercalatum.
S. haematobium
Fig. 12. Intravenous pyelogram of a woman infected with S. haematobium showing bilateral hydronephrosis
with deformity of both ureters.
factors, such as the presence of nitrosamines parts of the genital tract, including the
in the urine, are probably involved (Mostafa cervix, vagina and vulva, and about 6–27%
et al., 1995). Squamous carcinoma is more of such cases result in sterility.
common than transitional carcinoma (Fig.
13). It has been estimated that schistosomia-
S. japonicum, S. mekongi, S. mansoni
sis is responsible for about 16% of cases of
and S. intercalatum
bladder cancer in Egypt. In the Nile Delta
region, men do most of the agricultural
work and thus become infected, resulting in Invasive phase. As for S. haematobium.
a 12:1 male-to-female bladder cancer ratio.
Pulmonary arteritis progressing to irrevers- Acute phase. Allergic manifestations, such
ible and lethal cor pulmonale because of as pyrexia, headache, oedema, cough,
capillary damage by eggs sometimes occurs dysenteric symptoms, pruritus and
when eggs are swept back into the lungs. urticaria, occur 3–8 weeks after infection
The presence of adult worms in the lungs with S. japonicum. This is known as the
following drug treatment is also a possible Katayama syndrome and may be accompa-
cause of pulmonary damage. In women, nied by tenderness in the liver region, mild
eggs may cause lesions in the ovaries, abdominal pain, lymphadenopathy and
Fallopian tubes and uterus or in the lower splenomegaly, with an accompanying
20 Chapter 1
Fig. 13. Section of bladder. Eggs of S. haematobium (many are calcified or ‘black’ eggs) can be seen
with epithelial squamous cell metaplasia (arrowed).
Fig. 14. Egg granuloma of S. mansoni in liver with surrounding epitheloid cells and some leucocytic
infiltration (see Plate 1 for more advanced stage).
Fig. 15. Two boys with advanced schistosomiasis mansoni. Note collateral venous circulation in nearest
patient.
lobules. Liver enlargement is common and Sahara and signs of portal hypertension are
splenomegaly often follows the portal hyper- always present in such cases. Anaemia may
tension (Fig. 15). Hepatosplenic schisto- be found when there is splenomegaly and is
somiasis mansoni is more common in more severe after repeated haematemesis.
Brazil and Egypt than in Africa south of the The reaction to the eggs in the liver may
22 Chapter 1
Fig. 16. ‘Symmer’s clay pipestem’ fibrosis caused by eggs of S. japonicum surrounding portal veins in liver.
eventually cause the periportal fibrotic distributed throughout the body, and the
reaction termed ‘Symmer’s clay pipestem’ greater longevity of the adult worms.
fibrosis (Fig. 16). Liver function tests, how- Brain involvement is most common in S.
ever, are not altered in schistosome fibrosis, japonicum infections and two types of brain
as they are in true cirrhosis, and, although lesions have been reported. In the first type
there may be severe pathological lesions, there is diffuse involvement with scattered
there is no liver failure. Severe disease, lesions, probably caused by eggs being car-
with hepatosplenomegaly, occurs in about ried to the brain in the bloodstream, and
10% of cases of schistosomiasis mansoni, this type usually results in no symptoms. In
but takes 5–15 years to develop, and in the second type a localized granulomatous
children infection can have effects on nutri- mass is present, containing large numbers of
tion (Stephenson, 1993). eggs deposited by ectopic adult worms in
As the portal pressure increases, a the blood-vessels of the brain. These granu-
collateral venous circulation becomes lomas can cause a wide range of symptoms,
established and severe or even fatal depending on the anatomical location of the
episodes of bleeding can result from the lesion in the brain.
oesophageal varices. The portal systemic A transverse myelitis can result from the
shunt results in the eggs bypassing the presence of eggs in the spinal cord and is
liver and being deposited in the lungs. An most commonly found in infections with S.
obstructive and destructive arteritis may mansoni and S. haematobium.
follow, which can lead to systematic arter- In order for the eggs to make their way
ial hypertension and eventually to hyper- through the tissues into the gut, the
trophy of the right ventricle. miracidia release proteolytic enzymes and
The adult worms in the blood-vessels do other material, which diffuses through
little damage when living but their death pores in the eggshell. This material (solu-
can lead to focal necrosis of the liver cells ble egg antigen (SEA)) is highly immuno-
and to granulomas. The more severe genic, and the immune response made
pathology thought to be produced by infec- against it leads to the formation of the large
tion with S. japonicum is usually granulomas that are responsible for most of
explained by the greater egg production, the pathology in this phase. Evidence for
the spherical shape of the egg lacking a this and analysis of the mechanisms involved
large spine resulting in more eggs being have come largely from experiments in
The Trematodes 23
mice. Granulomas can be induced in the occur against these species (Wilkins et al.,
lungs of mice by intravenous injection of 1984, 1987; Butterworth, 1998; Kabatereine
eggs, and this process is prevented by et al., 1999), and studies in Brazil link resis-
immunosuppressive treatments that inter- tance to particular genetic characteristics
fere with cellular responses. The degree of (Abel and Dessein, 1997). Infections with S.
response to eggs and the pathology that japonicum do not show this picture,
results, following infection in mice, are although there appears to be an age-related
strongly influenced by genetic factors, and reduction in pathology (Ross et al., 2000).
this also reflects different degrees of The slow acquisition of immunity against S.
immune responsiveness. It has recently haematobium and S. mansoni correlates
been shown in humans that at least some with a change in the balance between
of the variation in pathology seen in antiparasite immunoglobulin E (IgE) and
infected populations is due to the activity IgG4, the former promoting protective
of particular major genes, which are associ- responses against incoming larvae, perhaps
ated with T-cell function (e.g. a codomi- by antibody-dependent cellular cytotoxic
nant major gene, SM1, on chromosome 5). (ADCC) mechanisms directed against anti-
The immunology of granuloma formation gens expressed on the surface tegument. As
is complex and involves the activity of shown by work carried out in the 1960s,
both major subsets of CD4+ T-helper (Th) adult schistosomes are largely unaffected by
cells. SEA is a powerful inducer of Th2 immunity and continue to survive in hosts
responses, but Th1 cells also play a role. immune to larval stages (concomitant immu-
The cellular response is initiated and con- nity). One way in which this is achieved is
trolled by the cytokines released and in by the adult schistosomes masking the for-
mice the phenomenon of immunomodula- eign nature of their surface antigens by
tion occurs – i.e. early granulomas tend to incorporating host-derived molecules into
be much larger than those formed later in the tegument and by rapid replacement of
infection. the tegument (antigenic disguise).
Antigen–antibody complexes have been Schistosomes, like many other helminths,
shown by fluorescent antibody studies to exert profound effects on the host’s immune
be the cause of the Splendore–Hoeppli response. Not only do these parasites tend to
phenomenon that occurs in sensitized polarize the T-cell response selectively
hosts. Glomerulonephritis has been towards Th2 activity, but they may also
reported as an immune-complex disease in modulate other components in ways that
schistosomiasis mansoni. result in a more general suppression of
immune and inflammatory responses.
IMMUNOLOGY OF SCHISTOSOMES Immunomodulation can influence the
Although it has been established for many ability of the host to respond successfully
years that laboratory animals, including pri- to other infections, and there is recent evi-
mates, develop immunologically mediated dence that schistosomes and soil-transmit-
resistance to experimental infections, defi- ted nematodes may increase susceptibility
nite proof that acquired immunity to schisto- to both AIDS and tuberculosis (Bundy et
some infections occurs in humans has been al., 2000).
very difficult to demonstrate. It is clear that
intensity of infection and prevalence with S.
DIAGNOSIS
haematobium and S. mansoni is greatest in
10–14-year-olds and declines in older age- Parasitological. The presence of eggs in the
groups, but this could be explained by a faeces or urine is still the most widely used
decrease in water contact. However, longitu- method of diagnosis. This usually poses no
dinal studies and detailed monitoring of problems in heavy infections but diagnosis
reinfection after elimination of an existing may not be so easy in light infections, as
infection by chemotherapy have demon- occur in the majority of cases, particularly
strated that age-dependent resistance does in tourists, who have probably been infected
24 Chapter 1
only once. In general, it is only the presence each species more accurate estimates are
of eggs containing live miracidia that possible (Guyatt et al., 1999).
indicates an active infection requiring treat- Portable ultrasound can be used for
ment. diagnosis of the degree of pathology, partic-
The eggs of S. haematobium, which occur ularly in the liver or bladder, and can
in the urine, can be detected by sedimenta- screen populations at the community level;
tion of a 10 ml sample (best collected around it can also be used for determining the
midday) in a urinalysis flask, the deposit effect of chemotherapy (Hatz, 2000).
being examined under the microscope for
eggs. Alternatively, a 10 ml sample can be Immunological. There are numerous
passed through a polycarbonate or poly- immunological tests for diagnosis but those
amide membrane, washed, removed and involving various ELISA and immunoblot-
stained in 1% trypan blue (or 2.5% ninhy- ting techniques are the most convenient. A
drin, 5% iodine, eosin or methylene blue) dipstick ELISA for urine samples, using an
for a few minutes. After washing again (and SEA can effectively diagnose schistosomia-
drying at 37°C if wished), the filter is exam- sis and correlates well with quantitative
ined in a few drops of saline on a micro- egg counts. Circulating cathodic antigen
scopic slide. It is probably not wise to reuse (CCA) is the dominant antigen in the urine
filters. In about 5% of patients, eggs may also of S. mansoni patients and can be detected
be found in the faeces. by a monoclonal antibody (mAb) of 41/42
Cytoscopy may be performed as a final kDa. There are also circulating soluble
diagnostic measure when no eggs can be adult-derived antigens (Sm31/32 and
found in the urine, but the danger of Sj31/32) in serum, which can be detected
secondary infection makes this a possibly by dot ELISA, and an IgG ELISA can be
hazardous procedure. used to detect antibodies against them.
The eggs of the other species occur in
the faeces and concentration techniques, TREATMENT
such as the formol–ether or Kato–Katz Chemotherapy. Praziquantel (usually in a
thick-smear methods (p. 258), are necessary. single oral dose of 40 mg kg1) is the drug of
A modification of the thick-smear method choice for all species of schistosome and has
can be used for quantitative determinations virtually superseded almost all the agents
and, in some studies, has given a good previously used. It has few side-effects
indication of the number of adult worms, (except sometimes in patients with very
as can filtration staining techniques. In the heavy worm loads), although it is probably
latter, eggs from faeces are stained with better not to give it during early pregnancy.
ninhydrin on filter-paper and counted. Praziquantel is a heterocyclic pyrazine-
Rectal biopsy is often effective (even in isoquinoline unrelated to other
about 50% of cases of S. haematobium), an anthelminthics – chemical formula 2-(cyclo-
unstained squash being examined under hexylcarbonyl)-1,2,3,6,7,11-b-hexahydro-4H-
the microscope. pyrazino[2,1a]isoquinolin-4-one. It is quickly
metabolized, crosses the blood– brain barrier
Clinical. In the field, haematuria can provide and appears to act by causing spastic paraly-
a quick and inexpensive indicator of urinary sis and vacuolization of the tegument of the
schistosomiasis; microhaematuria (measured worm, possibly from interference with inor-
by means of a reagent strip) or proteinuria ganic ion transport and an increase in mem-
correlates well with the intensity of infec- brane permeability; it may also destroy
tion. Even self-reporting by children can lysosomes. Two target antigens are also
provide a useful epidemiological record of affected (a surface tubercle glycoprotein of
community prevalence, although this is 200 kDa and an esterase of 27 kDa) and
likely to give an overestimate, depending on maybe epitopes become exposed so that the
the species of worm and level of infection drug evokes an effective immune response
(Ansell and Guyatt, 1999); using models for (Brindley, 1994; Dupre et al., 1999).
The Trematodes 25
*It is probable that in these countries transmission has now ceased and these are residual cases.
26 Chapter 1
Gabon (29% in one area), São Tomé and munity that has levels of egg output likely
Principe 0.005 (4%). Confirmation needed to cause morbidity (Jordan et al., 1993).
for presence in Central African Republic, This demonstrates the importance of quan-
Chad, Mali and Nigeria. titative determinations of egg output in
schistosomiasis (p. 260). The ecological fac-
S. mekongi. Cambodia 0.07 (0.7%), Laos tors associated with the transmission of
0.12 (2%) and recent cases in refugee schistosomiasis vary markedly with the
camps in Thailand. species of schistosome, owing to the differ-
ing habitats of the snails involved. The
S. malayensis. Malaysia very rare (Shekar, amphibious snail hosts of S. japonicum live
1991). mainly in rice paddies and muddy habitats
beside rivers; the aquatic snail hosts of S.
In most areas endemic for S. haematobium mansoni live principally in gently flowing
there is a very high prevalence rate, with running water (less than 0.3 m s1 ) such as
almost everyone in the community lightly irrigation channels, streams, lakes and
infected before the age of 10 years. Studies ponds (Plate 2); those transmitting S.
in Brazil, St Lucia, Tanzania and Uganda haematobium live almost entirely in still
demonstrated a relationship between egg water, such as ponds, pools, lakes and
output and prevalence of S. mansoni in marshy areas. Snail populations usually
children (Fig. 17). Above 50–60% preva- show marked seasonal fluctuations, the
lence, a small extra rise is accompanied by most important factors influencing numbers
a considerable increase in egg output and being rainfall and temperature. The effect
thus in the severity of disease. Post-mortem of rainfall varies, depending on the yearly
studies and blood-filtration studies after rainfall figures. In drier regions of Africa
chemotherapy for counting adults have rainfall stimulates the production of young
shown a good relationship between the last bulinid and planorbid snails, while in
two parameters so that intensity of infec- equatorial regions with a higher rainfall,
tion can be expressed in terms of eggs per snails are washed away during the rainy
gram of faeces or per 10 ml of urine. To season and breeding takes place principally
determine the intensity of infection of dif- in the months following. Temperature is
ferent age-groups, the frequency of occur- another factor limiting population densi-
rence of the different levels of egg output ties, the temperature range for optimal
can be conveniently plotted and provides a expansion of snail numbers being
good indicator of the proportion of the com- 20–28°C.
100 x ZAIRE
x KENYA
Prevalence (%)
75 x EGYPT
x BRAZIL
x ST LUCIA x ZAMBIA
50 x KENYA
x ETHIOPIA
x BURUNDI
x BURUNDI
25
x ETHIOPIA
Fig. 17. Relationship between overall prevalence and intensity of S. mansoni infection as determined by
egg output (measured by Kato–Katz technique) in various endemic areas. Each determination is based
on studies by different workers. e.p.g., eggs per gram. (From Jordan et al., 1993.)
The Trematodes 27
Seasonal variations in the cercarial trans- hemisphere B. glabrata is the most impor-
mission rates in snails are most marked with tant snail host, with seven other species
the bulinid hosts of S. haematobium living locally involved.
in temporary bodies of water. In hyper-
endemic habitats, such as canals, there may S. haematobium. Bulinus is a turreted pul-
be large numbers of snails producing cer- monate snail with a left-handed opening
cariae of S. mansoni throughout the year. when looked at with the spire upwards
Diurnal fluctuations in the production (Fig. 1). The height varies from 4 to 23 mm.
of cercariae occur; cercariae of S. japon- The subgenus B. (Physopsis) is differenti-
icum (which is adapted for transmission ated from B. (Bulinus) by having a truncate
among the nocturnal animal reservoir columella and a pointed end to the foot in
hosts) are mostly produced in the evening, living specimens. Two main groups of
while those of S. mansoni and S. haemato- bulinid snails are important as hosts: (i) the
bium are produced in the middle of the africanus group (ten species) of the sub-
day. This factor is not usually of practical genus Physopsis is the more important
importance except where S. mansoni trans- group in eastern and southern Africa and
mission occurs in running water, in which in most parts south of the Sahara; and (ii)
case washing and bathing are likely to be the truncatus, forskali and reticulatus
much safer in the morning and evening, groups (27 species) of the Bulinus sub-
when cercariae are less abundant. genus of snails act as hosts in the Near East
(Iran, Egypt and Sudan), in Madagascar
SNAIL HOSTS OF SCHISTOSOMES and in parts of both East and West Africa.
S. japonicum. Oncomelania is a conical,
amphibious, prosobranch snail measuring S. intercalatum. B. (Physopsis) africanus acts
3–10 mm and having 4–8 whorls, a dextral as snail intermediate host in Congo Republic
opening and an operculum covering the foot. and B. (Bulinus) camerunensis in Cameroon.
Species involved are O. hupensis from main- It must be borne in mind when attempting
land China, O. nosophora from south-west to identify a snail that there are usually
China (and Japan), O. formosana from many other types of snail present in the
Taiwan, O. quadrasi from the Philippines same body of water as the ones transmit-
and O. lindoensis from Sulawesi. ting schistosomiasis, and they may be
difficult to differentiate. Specialized mono-
S. mekongi. Neotricula aperta is an aquatic graphs need to be consulted (Brown, 1980)
snail found in the Mekong River and it or expert advice sought. It is also important
probably also transmits S. sinensium. to distinguish clearly the cercarial species
Robertsiella spp. are found in Malaysia and emerging, because of the large number of
transmit S. malayensis. animal and bird schistosomes that use the
same snail hosts and produce cercariae
S. mansoni. Biomphalaria is a flattened pul- similar to the human schistosomes. In
monate, planorbid snail with 3.5–7 whorls, detailed epidemiological studies, it may be
measuring 7–22 mm. Four main groups are necessary to expose laboratory animals to
found in Africa: (i) the pfeifferi group (four the cercariae and to identifiy any adult
species), which include the most important schistosomes that develop.
hosts in Africa and the Middle East; (ii) the
sudanica group (three species), which occur
in both East and West Africa; (iii) the PREVENTION AND CONTROL
choanomphala group (three species), which Personal prevention is by avoidance of
live in the great lakes and act as hosts in water sources containing cercariae, impos-
Lake Victoria; and (iv) the alexandria group sible for fishermen to carry out and
(two species), which occur sporadically in extremely difficult for children to be con-
North, East and South Africa. In the western vinced of the necessity thereof. The most
28 Chapter 1
or 0.2 g m2 on moist soil for amphibious avoiding contaminated water needs to be
hosts of S. japonicum. effectively communicated, particularly to
children and adolescents. While it is prob-
Biological control. Carnivorous snails, such ably impossible in Africa to prevent chil-
as Marisa cornuarietis, or competitors, dren swimming in the water, they could be
such as Melanoides tuberculata and taught to urinate (and perhaps defecate)
Tarebia granifera, have been added to habi- before entry.
tats in Guadeloupe, Martinique, Puerto
Rico and St Lucia, with the result that Present position and future outlook for con-
Biomphalaria has vanished from all or trol. Present control methods are capable of
almost all areas (Giboda et al., 1997; substantially reducing transmission and
Pointier and Jourdane, 2000). particularly morbidity in most areas pro-
vided that they are properly designed and
Ecological methods of control. Habitats can carried out and if adequate funds are avail-
be made unsuitable for snails by alternate able. However, new irrigation and hydro-
flooding and drying of water channels or electric schemes in many endemic
covering and lining of canal systems, as in countries have increased the number of
Egypt and Sudan, and filling in of marshy snail habitats. New human-made lakes
areas. In many areas such measures are have followed the building of dams in
likely to be permanently successful, but many countries: Côte d’Ivoire (Lakes
they require close cooperation between Kossou and Taasbo), Egypt and Sudan
irrigation engineers and public health (Lake Nasser); Ghana (Volta Lake); Nigeria
workers. In rice-growing areas of mainland (Kainji Lake); Zambia and Zimbabwe (Lake
China and in Leyte Island (Philippines) Kariba); Senegal (Lake Guiers below the
widespread cleaning of irrigation ditches Diama dam and Manantali dam in Mali).
and filling in of ponds have resulted in a The incidence of schistosomiasis in the
great reduction in schistosomiasis japon- populations bordering these lakes or the
icum infection in recent years. irrigation channels coming from them (as
in Egypt and the Gezira scheme in Sudan)
Prevention of water contact. The provision almost always rises. In Côte d’Ivoire
of piped water supplies and alternative around Lake Kossou there used to be an
bathing and clothes-washing places can be infection rate of 14% with S. haemato-
effective in preventing human contact with bium, which has now risen to 53%, and
infested water and, while expensive, does around Lake Taabo the rate has risen from
have wide-ranging health benefits. 0% to 73% (although the 2% infection rate
For visitors and tourists, prevention can with S. mansoni has not changed so far);
be achieved by avoidance of contact with around Lake Kainji the infection rate with
water in ponds, canals, slow-flowing S. haematobium is now 14% in children
streams and the shallow edge of lakes in (from nothing); around Lake Kariba there is
endemic areas (although in Lake Victoria in now a 70% infection rate in children with
East Africa the snail Biomphalaria S. haematobium; infection has spread
choanomphala transmits S. mansoni in greatly in the Senegal River basin since the
water 2 m deep). For tourists, swimming in building of the dams and there is now a
Lake Malawi or Lake Kariba is often fol- prevalence of 91% with S. mansoni (from
lowed by infection. Dibutyl phthalate or 1%) and of 28% with S. haematobium
hexachlorophene spread over the skin will (Southgate, 1997). One trend in Africa
protect for about 4 h. caused by changes in ecology following the
building of dams in Africa (e.g. in delta
Health education. Knowledge of the role of areas of Cameroon, Ghana and Egypt) is the
indiscriminate defecation and urination in replacement of S. haematobium by S. man-
spreading the disease to members of a com- soni. The new opportunities for agriculture
munity and the personal importance of are also attracting possibly infected
30 Chapter 1
migrants and this is particularly apparent Progress to date indicates that a vaccine is
in the Gezira and in Ethiopia (although the feasible, since protection against sheep
building of the Koka dam initially reduced schistosomiasis (a problem in Sudan) using
transmission). irradiated cercariae has been successfully
Infection has been eliminated from achieved. However, any human vaccine
Japan, Tunisia and Montserrat recently and will have to consist of defined components,
campaigns against S. haematobium are as cercariae might introduce viruses.
being mounted in Mauritius (1.3% in adults Various types of vaccine are being inves-
only), Morocco (2.3% overall) and Tanzania tigated (Capron, 1998). These include the
(45% in Zanzibar), against S. mansoni in following.
Brazil, Burundi, St Lucia and Venezuela
1. Peptide components of protective anti-
(risen from 0.1% to 0.9% in under-10-year-
gens. These can be used to generate
olds in the last few years) and against both
differential responses, e.g. one elicits
species in Egypt, Ghana, Madagascar,
eosinophil-dependent ADCC, another stim-
Malawi, Mali, Nigeria, Saudi Arabia, Sudan
ulates delayed-type hypersensitivity (DTH)
and Zimbabwe; against S. japonicum in
reactions with macrophage activation.
China, Indonesia (1% overall infection rate)
and the Philippines (3.6%); and against S. Recently a multiple antigen peptide (MAP)
mekongi in Laos (72%) (WHO, 1993). vaccine has shown very promising results,
China has had control campaigns involv- inducing both T-cell and B-cell responses
ing mass treatment of people and cattle with in mice and in monkeys, reducing parasite
anthelminthics (nowadays praziquantel), fecundity and egg viability and decreasing
snail control and environmental engineering liver pathology by over 70%.
for the last 50 years but there were still esti- 2. Excretory/secretory (ES) antigens. ES
mated to be 1.06 million cases in humans material obtained from postcercarial schisto-
and 0.25 million cases in cattle in 1997. somula kept in vitro is effective in immu-
However, S. japonicum has been eliminated nizing rats to subsequent infection. The
from 150 of the formerly 378 endemic coun- secretions contained proteins in the range
ties in 12 provinces south of the Yangtze 22–26 kDa (amplified by PCR) and induce
River (Ross et al., 1997), while in 1955 there an IgE-dependent ADCC.
were estimated to be over 11 million cases. 3. Membrane antigens. Molecules with a
Unfortunately, recent flooding has increased molecular weight of 23 kDa (e.g. Sm23 and
the extent of snail habitats. The Three Sj23) occur in the tegument of all stages of
Gorges dam under construction on the the parasite but are often species-specific.
Yangtze River will, within the next 10 years, 4. Paramyosin. This 97 kDa internal anti-
create the largest lake in the world in a gen (e.g. Sm97), obtained from schisto-
highly populous region where schistosomia- somula or adults, induces responses that
sis japonicum already occurs and where affect the muscles and tegument of adults
there will have to be about 2 million extra (Gobert, 1998).
migrants from highly endemic areas, which 5. DNA vaccines. Direct injection of the
will be flooded. The building of the dam coding sequence for known antigens is
should reduce the overall density of snails being investigated. Such a vaccine can be
along the river banks but will provide many used alone or combined with chemother-
more suitable, contaminated, marshland apy, e.g. immunization with DNA for Sm28
habitats and an increased contact with water. glutathione-S-transferase (GST) together with
praziquantel prevented the formation of
Towards a vaccine. The need for a vaccine pathological lesions in mice (Dupre et al.,
against schistosomiasis is underscored by 1999).
the fact that, although chemotherapy with 6. Irradiated live cercarial vaccines.
praziquantel is very effective, in the Although these are not directly suitable for
absence of an effective immune response human use, they might provide useful
reinfection tends to occur quite quickly. pointers to promising leads (Coulson, 1997).
The Trematodes 31
The most promising six antigens were on the island of Leyte in the Philippines,
recently selected by the World Health although many other species of animals are
Organization (WHO) for testing as vaccines infected, as they are on Sulawesi (Indonesia)
but at best gave only a maximum of 60% and used to be in Japan. Molecular studies
protection (Katz, 1999). indicate that S. japonicum is a species com-
A general problem facing the develop- plex of many sibling species, and in Taiwan
ment of vaccines against any infectious dis- the local form of S. japonicum occurs only
ease is the generation of an appropriate in animals and is not infective to humans.
T-cell response (e.g. CD4+ or CD8) and, if The dog appears to be the only reservoir host
CD4+, then the appropriate Th subset, Th1 of S. mekongi in Laos. S. sinensium is a little
or Th2. In addition, where the disease asso- known species with a laterally spined egg
ciated with infection is immune-mediated, found a few times in humans and rodents in
as in schistosomiasis, vaccination must not China and Thailand (Greer et al., 1989),
induce pathological responses. There is while the ‘Tonle Sap schistosome’ has been
controversy about whether Th1 or Th2 newly discovered in humans living around
responses are most important and appro- Lake Tonle Sap in Cambodia (Pecoulas et al.,
priate for vaccine-induced resistance to 1995).
schistosome infection, but it is probable S. mansoni has been reported frequently
that a balanced response would be best from animals (of 38 species) but in many
(Wynn and Hoffmann, 2000). cases they do not appear to have any epi-
The route of administration of any vac- demiological significance. However, baboons
cine is also very important, as is the choice and green monkeys are apparently main-
of a suitable adjuvant. Presentation of anti- taining infections among themselves in
gens via an engineered vector organism can areas of East Africa and rodents may act as
overcome both problems. For example, reservoir hosts in East Africa, Senegal and
Bacillus Calmette–Guérin (BCG) has been Guadeloupe and particularly in Brazil,
engineered to produce an intracellular where up to 15 species appear to be of
schistosome GST and this has induced a increasing importance in maintaining
strong and long-lasting response in mice infection in urban and periurban areas of
when given by the intranasal route (a the north-east (Mott et al., 1995).
mixed response of GST-specific immuno- Animals appear to be of little or no
globulin – IgG2a, IgG2b and IgA in serum importance in the transmission of S.
and IgA in bronchial lavage fluids). A vac- haematobium, although various primates
cine against schistosomiasis haematobium have been found infected, nor are they for
(Sh28GST – Bilhvax) together with prazi- S. intercalatum.
quantel treatment is undergoing field trials The schistosomes found in humans and
in Senegal. animals in Africa can be divided into two
groups of sibling species. Hybridization
ZOONOTIC ASPECTS can occur experimentally within members
Animal hosts are extremely important in the of each group but is not common in nature.
transmission of S. japonicum and S. In the S. haematobium/S. intercalatum
mekongi and in some countries can maintain group, S. mattheei Veglia and Le Roux,
the parasite in the absence of humans. In 1929, is a parasite of sheep, cattle, wild
mainland China infection occurs in 40 animals and occasionally humans in south-
species of domestic and wild animals ern Africa. The eggs in the faeces or urine
(Jordan et al., 1993) and is important as a have a terminal spine and measure
serious cause of morbidity and mortality in 120–180 µm in length (Fig. 124). Curiously,
cattle and goats. In the Dongting Lake area of in all human cases, the eggs have occurred
Hunan, 60% of cattle and buffaloes, 24% of together with those of S. haematobium or
pigs, 9% of dogs and 7% of people were S. mansoni; the females of S. mattheei are
found to be infected. Dogs and pigs play an known to be capable of producing eggs
important part in maintaining transmission parthenogenetically and are perhaps carried
32 Chapter 1
33
34 Chapter 1
LIFE CYCLE the water, where they can survive for only
The eggs are usually swallowed and pass 24–48 h.
out in the faeces but may also escape in If cercariae find freshwater crabs, princi-
sputum. For further development they pally Eriocheir, the mitten crab in Asia and
have to reach water. The miracidia hatch also Potamon (Fig. 19) and Sesarma, or
after 3 weeks in water (at an optimum tem- freshwater crayfish (Astacus), they penetrate
perature of 27°C) and penetrate freshwater and encyst in the gills or muscles as meta-
snails. These are operculate genera, of cercariae (measuring 250–500 µm).
which the most important is Semisulcospira Freshwater crustaceans can probably also
(S. libertina is an intermediate host species become infected by ingesting unencysted
in China, Japan, Taiwan and Korea). Other cercariae in the water or even inside an
important snails are S. amurensis, Thiara infected snail.
granifera and Oncomelania nosophora (see The crustacean intermediate hosts are
Fig. 1). eaten by humans as food. If they are eaten
In the snail the miracidia develop into raw, the metacercariae hatch in the duode-
sporocysts followed by two generations of num and young worms penetrate through
rediae, and finally, in about 3 months, give the intestinal wall and pass across to the
rise to very short-tailed cercariae. The cer- abdominal wall in about 6 h. The immature
cariae emerge from the snail and swim in flukes burrow through the diaphragm to the
Fig. 18. Cut portion of lung (of a tiger) with cysts containing pairs of adult Paragonimus. Note large
excretory bladder in each fluke.
pleural cavity in 6–10 days and enter the become surrounded by a cyst, possibly
lung capsule in 15–20 days. The first eggs caused by softening followed by fibrosis
are passed 60–70 days after infection. (Plate 4). However, as the cyst wall is usu-
Although Paragonimus is hermaphrodite, ally lined with columnar epithelium, it is
two worms are necessary for fertilization to probable that, in most cases, it represents
take place (Fig. 19 and Plate 4). This is true the expanded wall of a bronchus. The cap-
for the usual diploid form of P. westermani, sule is collagenous and oedematous, with
but there is also a triploid form, in which plasma cells, eosinophils, neutrophils,
parthenogenesis can occur; the adults of the macrophages and fibroblasts. The cyst mea-
latter are larger and more pathogenic and sures about 1 cm in diameter after 90 days
the eggs differ in size and shape (see Blair and is filled with a thick brownish fluid
et al., 1999). The adult worms can live for containing eggs. Bronchopneumonia may
20 years but usually die after about 6 years. result when a cyst bursts or retains an
opening into a bronchus. Eggs are often
CLINICAL MANIFESTATIONS present in the lung tissues and cause
There are no recognizable symptoms pseudotubercles similar to those caused by
accompanying the migratory phases. schistosome eggs, with infiltration by
The first signs are usually fever with a eosinophils and lymphocytes, followed by
dry cough, sometimes accompanied by giant cells and fibroblasts.
bloodstained sputum containing eggs after
rupture of the cysts. Chest pain can be Extrapulmonary. Adults are sometimes
severe and, if the infection is heavy, there found in many parts of the body, particu-
is sometimes increasing dyspnoea and larly in the organs of the abdominal cavity
bronchitis. Many patients with haemo- or subcutaneous tissues. In these sites the
ptysis are diagnosed as suffering from worms are rarely fertilized and presumably
pulmonary tuberculosis. have developed from larvae that lost their
Cerebral paragonimiasis may result in way. This occurs more frequently with
epileptic seizures, as well as headaches, species of Paragonimus which are less
visual disturbances and symptoms of adapted to humans than P. westermani.
meningitis. Most serious complications follow the
presence of flukes in the brain (Fig. 20).
PATHOGENESIS
This may be common in areas of high
Pulmonary. As flukes grow in the lungs endemicity (e.g. over 5000 cases per
they cause an inflammatory reaction and annum in South Korea). Adults found in
Fig. 20. Radiograph of head of a girl with soap-bubble type of calcification surrounding cysts containing
adult Paragonimus in occipital lobes.
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“The hounds bay and the hunt sweeps off in the distance—now lost to sight, and
now emerging from the hollows. The volunteers soon begin to return, and are seen
everywhere straggling about over the slopes. The carriages move on,
accompanying, as they can, the hunt by the road, till it strikes across the country
and is lost. The sunshine beats on the mountains that quiver in soft purple; larks
sing in the air; Brown, Jones, and Robinson ride by the side of the carriages as they
return, and Count Silinini smiles, talks beautiful Italian, and says, ‘Yas.’ He is a
guardia nobile, and comes to the house twice a week if there are no balls, and
dances with Marianne at all the little hops. Signor Somarino pays his court
meanwhile to Maria, who calls him Prince, emphasising the title when she meets
her friends the Goony Browns. And so the hunting picnic comes back to Rome.”
“‘Eh! Speriamo’ (let us hope so), was the answer of the pifferaro, as he showed
all his teeth in the broadest of smiles. Then, with a motion of his hand, he set both
the young men going, he himself joining in, straining out his cheeks, blowing all
the breath of his body into the little pipe, and running up and down the vents with
a sliding finger, until finally he brought up against a high, shrill note, to which he
gave the full force of his lungs, and after holding it in loud blast for a moment,
startled us by breaking off, without gradation, into a silence as sudden as if the
music had snapped short off like a pipe-stem.”
“My friend Count Cignale is a painter—he has a wonderful eye for colour and an
exquisite taste. He was making me a visit the other day, and in strolling about the
neighbourhood we were charmed with an old stone wall of as many colours as
Joseph’s coat: tender greys, dashed with creamy yellows and golden greens and
rich subdued reds, were mingled together in its plastered stonework; above
towered a row of glowing oleanders covered with clusters of roseate blossoms.
Nothing would do but that he must paint it, and so secure it at once for his
portfolio; for who knows, said he, that the owner will not take it into his head to
whitewash it next week, and ruin it? So he painted it, and a beautiful picture it
made. Within a week the owner made a call on us. He had seen Cignale painting
his wall with surprise, and deemed an apology necessary. ‘I am truly sorry,’ he said,
‘that the wall is left in such a condition. It ought to be painted all over with a
uniform tint, and I will do it at once. I have long had this intention, and I will no
longer omit to carry it into effect.’
“‘Let us beseech you,’ we both cried at once, ‘caro conte mio, to do no such thing,
for you will ruin your wall. What! whitewash it over!—it is profanation, sacrilege,
murder, and arson.’
“He opened his eyes. ‘Ah! I did not mean to whitewash it, but to wash it over
with a pearl colour,’ he answered.
“‘Whatever you do to it you will spoil it. Pray let it alone. It is beautiful now.’
“‘Is it, indeed?’ he cried. ‘Well, I hadn’t the least idea of that. But if you say so, I
will let it alone.’
“And thus we saved a wall.”
The preceding scrap reminds us of a passage from Alphonse Karr,
one of the most quietly-humorous of living French writers, who
relates, in one of his quaint, dreamy, desultory books, how a
neighbour of his, who lived in a poor thatched cottage on the fringe
of a wood, embowered in flowers, shaded by venerable trees,
refreshed by the balmiest of breezes, and enlivened by the songs of
countless birds, suddenly disappeared from the countryside. Karr,
who had long admired the sylvan retreat, and almost envied its
occupant, inquired his fate. He had become rich, he was told; a
legacy had enabled him to go and live in the town. He could afford to
rent two rooms with new furniture and a gaudy paper, and he looked
out upon a dirty street, along which omnibuses continually rolled.
“Poor rich man!” Karr pitying exclaims. He had whitewashed his
wall.
The Roman Ghetto furnishes the theme of one of Mr Story’s
longest and most lively chapters; Fountains and Aqueducts, Saints
and Superstitions, the Evil Eye, are the titles of three others. He
begins his second volume with a vivid and characteristic sketch of the
Markets of Rome, which are well worth the attention of foreign
visitors, especially of Englishmen, who will find their arrangements,
and much of what is there sold, to contrast strikingly with what they
are accustomed to in their own country. Carcasses of pigs and goats
adorned with scraps of gold-leaf and tinsel, blood puddings of a
brilliant crimson, poultry sold by retail—that is to say, piecemeal, so
that you may buy a wing, a leg, or even the head or gizzard of a fowl,
if so it please you. There is game of all sorts, and queer beasts and
fowls of many kinds are also there; the wild boar rough and snarling
—the slender tawny deer—porcupines (commonly eaten in Rome)—
most of our English game-birds—ortolans, beccaficoes, and a great
variety of singing-birds. Passing into the fruit and vegetable market,
one comes upon mushrooms of many colours, and some of them of
enormous size, most of which would in England be looked upon as
sudden death to the consumer, although in Italy they are found both
savoury and harmless. “Here are the grey porcini, the foliated
alberetti, and the orange-hued ovole; some of the latter of enormous
size, big enough to shelter a thousand fairies under their smooth and
painted domes. In each of these is a cleft stick, bearing a card from
the inspector of the market, granting permission to sell; for
mushrooms have proved fatal to so many cardinals, to say nothing of
popes and people, that they are naturally looked upon with
suspicion, and must all be officially examined to prevent accidents.”
Besides the fruits common in England, figs are very abundant, and of
many kinds; and when the good ones come in, in September, the
Romans of the lower classes assemble in the evenings, in the Piazza
Navona, for great feeds upon them. Five or six persons surround a
great basket and eat it empty, correcting possible evil results by a
glass of strong waters or a flask of red wine. But figs are a wholesome
fruit—much more so than one which at Rome, and in many parts of
Southern Europe, is the most popular of all—namely, the water-
melon. What millions of people, from the Danube’s banks to the
Portuguese coast, are daily refreshed the summer through by those
huge green gourds, hard and unpromising in outward aspect, but
revealing, at stroke of knife, rich store of rosy pulp, dotted with sable
seeds! Pesth is a great place for them; and daily, when morning
breaks, so long as they are in season, they are to be seen piled, all
along the river-side, in heaps like those of shot and shell in an
arsenal, only much broader and higher. All through the hot months,
in Hungary’s pleasant and interesting capital, few persons think of
dining without associating with the more heating viands a moiety or
enormous segment of one of those great cold fruits—a strange
digestive, as we Northerners should consider it, but found to answer
well in sultry climes. At Rome they are equally appreciated, and are
set above the choicest grapes. People make parties to go out of the
city and eat them; and this was especially the case some years ago,
when the authorities forbade their entrance on account of the
cholera, but were unable to prevent their extramural consumption.
In ordinary times you find heaps of them in the streets, especially in
the Piazza Navona, that great mart of fruit and frippery, vegetables,
old books, brilliant handkerchiefs, and other finery for the market-
women—old iron, old bottles, and rubbish of all kinds—amongst
which miscellany the patient investigator may sometimes discover
valuable copies of the classic authors and precious antique intagli, to
be purchased for a mere song. Here, as the story goes, a poor priest
once bought, for a few baiocchi, a large cut-glass bead which took his
fancy, and which a friend, more knowing than himself, afterwards
discovered to be a diamond of great value, now belonging, we are
told, to the Emperor of Russia. The priest disappeared, which leaves
any ingenious and inventive writer full liberty to build a romantic
tale upon the incident. The natural finale of the affair, Mr Story
opines, would have been for the priest to have married the Emperor’s
daughter, but his being in orders was an impediment; and so we are
justified in presuming that some less agreeable means was found of
easing him of his jewel, which, when he first possessed it, he took to
be a drop from a chandelier, but to which he of course clung with
desperate tenacity when enlightened as to the quality of the gem.
Rome ought to be a good preserve for fiction-writers, there are so
many family histories, traditions, and anecdotes current there, which
would serve the novelist’s turn. Edmund About availed himself of
one such in his tale of ‘Tolla;’ and another over-true tale was
interwoven, not very long since, in a pleasant novelet of Roman life
in the pages of this Magazine. Mr Story’s volumes abound in
suggestive passages of the kind. If Rome be an admirable residence
for an artist (and for some of the reasons why it is so, see the ‘Roba,’
i. p. 66, 67), it ought also to be an excellent one for a writer, were it
not that it is found by many unfavourable to mental exertion. This is
said to be particularly exemplified in the case of diplomatists, many
of whom, after a certain time passed in the Papal capital, are apt to
conceive an intense dislike to despatch-writing, and to keep their
Governments extremely uninformed concerning the state of the Holy
City and the prospects of Pontifical politics. We remember to have
been told, when in Rome, the names of more than one foreign
minister who had been recalled, it was asserted, for no other reason
but that nothing could induce him to write despatches. Rome is
certainly one of the places where there is most temptation, at least
for one half of the year, to neglect business for pleasure; but there is
possibly also something in the climate which disinclines many people
to headwork. It is much the fashion to abuse the Roman climate; and
this has been done, especially of late, by persons desirous to show
that Rome is an undesirable, because a highly insalubrious, capital
for united Italy. It is to be feared the grapes are sour, and that the
yellow flag now hoisted would be struck at the same time with the
French tricolour. Our own experience and observations induce us
very much to concur with those passages of Mr Story’s book which
relate to this question. “Rome has, with strangers, the reputation of
being unhealthy; but this opinion I cannot think well founded—to the
extent, at least, of the common belief.” Many maladies, virulent and
dangerous elsewhere, are very light in Rome; and for lung
complaints it is well known that people repair thither. The “Roman
fever,” as it is commonly called (intermittent and perniciosa), is
seldom suffered from by the better classes of Romans; and Mr Story
(who speaks with authority after his many years’ residence in Rome)
believes that, with a little prudence, it may easily be avoided. The
peasants of the Campagna are, it is well known, those who chiefly
suffer from it, and why? “Their food is poor, their habits careless,
their labour exhausting and performed in the sun, and they sleep
often on the bare ground or a little straw. And yet, despite the life
they lead and their various exposures, they are, for the most part, a
very strong and sturdy class.” Mr Story gives it as a fact that the
French soldiers who besieged Rome in ‘48, during the summer
months, suffered very little from fever, although sleeping out on the
Campagna; but they were better clothed and fed, and altogether
more careful of themselves, than the native peasants. Generally
speaking, the foreigners who visit Rome are less attentive than the
Romans to certain common rules for the preservation of health. They
eat and drink too much, and of the wrong things. They get hot, and
then plunge into cold churches or galleries; whereas an Italian flies
from a chill or current of air as from infection. Mr Story gives a few
simple rules, by following which he declares you may live twenty
years in Rome without a fever. He cautions Englishmen against
copious dinners, sherry and brandy, and his own countrymen against
the morning-dinner which they call a breakfast; and supplies other
useful hints and practical remarks. The subject is one which interests
many, and such are referred to the ‘Roba,’ i. p. 156–161, and to the
chapter on the Campagna, in which high authorities and ingenious
arguments are brought to prove that in old times it was not
insalubrious, and that in our own it need not be so. Population and
cultivation are perhaps all that are needed to render tracts healthy
that now are pestilential, but which assuredly were not so in the time
of the ancient Romans, since many of them, we know, were their
favourite sites for patrician villas. Much might be done by an
intelligent and active government, and especially by a good sanitary
commission. There was one clever gentleman who wrote that Rome
was ill fitted to be the capital of Italy on account of its deficiency in
buildings suitable for government offices! Where good reasons are
not to be found silly ones may be resorted to, but they of course only
weaken the cause they are intended to prop. And if it were to be
urged that all the worst plagues flesh is heir to, combine to render
Rome for the present impossible as capital of Italy, the most we
could admit, by way of compromise, and borrowing a well-known
answer, would be, “non tutti, ma Buona parte.”
CAXTONIANA:
A SERIES OF ESSAYS ON LIFE,
LITERATURE, AND MANNERS.
By the Author of ‘The Caxton Family.’
PART XV.
Death, the only player that no man can cheat, cut into his table,
and trumped the last card of his long suit.
In the more brilliant period of this amiable man-scorner’s social
career, once, and once only, he is said to have given way to anger.
One of his associates (I say designedly associates, not friends, out of
respect for his memory, since friendship is a virtue, and he therefore
denied its existence)—one of his associates, warmed perhaps into
literature by his own polite acquaintance with all that is laide in
belles lettres, wrote a comedy. The comedy was acted. My hero
honoured the performance by appearing in the author’s box. Leaning
forward so as to be seen of all men, he joined his hands in well-bred
applause of every abortive joke and grammatical solecism, till, in a
critical part of the play, there occurred a popular claptrap—a
something said in praise of virtue and condemnation of vice. The
gallery of course responded to the claptrap, expressing noisy
satisfaction at the only sentiment familiar to their comprehension
which they had hitherto heard. But my archetype of modern
misanthropy paused aghast, suspended
“The soft collision of applauding gloves,”
“That the thought first occurred to him during his residence at Cambridge,
therefore before he had completed his fifteenth year, we know upon the best
authority—his own statement to Dr Rawley. I believe it ought to be regarded as the
most important event of his life—the event which had a greater influence than any
other upon his character and future course. From that moment there was
awakened within his breast the appetite which cannot be satiated, and the passion
which cannot commit excess. From that moment he had a vocation which
employed and stimulated all the energies of his mind, gave a value to every vacant
interval of time, an interest and significance to every random thought and casual
accession of knowledge—an object to live for as wide as humanity, as immortal as
the human race—an idea to live in vast and lofty enough to fill the soul for ever
with religious and heroic aspirations. From that moment, though still subject to
interruptions, disappointments, errors, and regrets, he could never be without
either work or hope or consolation.”
But this young philosopher is son of Sir Nicholas Bacon, the late
Lord Chancellor; the Queen has laid her hand upon his head while
yet a boy, and called him her young Lord Keeper; he is nephew to the
Prime Minister; he dreams of courts, of place, of power. He must
unite his lofty speculations with the great affairs of State; he must
survey human knowledge from the high places of society. He enters
Gray’s Inn, is a student of the law, and his heart aches after office
and promotion.
There is one person very intimately connected with Bacon, whom
Mr Spedding has brought before us with a novel distinctness—his
mother, Lady Bacon. We are not aware that her presence will throw
much light on the character of her son, but henceforth, we are sure,
no biography of the son will be written in which this lady will not be
a conspicuous figure. She is one of those strongly-marked characters
that always please the imagination; dogmatic, perverse, full of
maternal anxiety, pious and splenetic, with marvellous shrewd sense
and a very ungovernable temper. The knowledge of her character
would enable us to answer one question. Presuming that any one
should think fit to ask why Bacon did not seek the retirement of
Gorhambury, the answer is quite ready. There would have been no
peace for him under the roof of his lady mother. Puritan and
termagant, his philosophy would have been “suspect” to her; and his
retirement would have been certainly denounced as unpardonable
sloth. She is a learned lady, mingles scraps of Latin and Greek in her
epistles, and she can write, when the occasion demands, in a very
stately English style—stately, but straightforward withal. Her son’s
epistolary style is often involved and verbose. He does not often
come so directly to the point as Lady Bacon does in the following
letter, written to Lord Burghley, in the interest of the Nonconformist
clergy, or Preachers, as they were then called. In a conference which
had lately taken place at Lambeth between them and the bishops, she
thinks they had not fair-play; she appeals, in their name, to her
Majesty and the Council:—
“They would most humbly crave, both of God in heaven, whose cause it is, and of
their Majesty, their most excellent sovereign here on earth, that they might obtain
quiet and convenient audience rather before her Majesty herself, whose heart is in
God his hand to touch and to turn, or before your Honours of the Council, whose
wisdom they greatly reverence; and if they cannot strongly prove before you out of
the word of God that reformation which they so long have called and cried for to be
according to Christ his own ordinance, then to let them be rejected with shame out
of the Church for ever.... And therefore, for such weighty conference they appeal to
her Majesty and her honourable wise Council, whom God has placed in highest
authority for the advancement of His kingdom; and refuse the bishops for judges,
who are parties partial in their own defence, because they seek more worldly
ambition than the glory of Jesus Christ.”
Soon after this Anthony does return home, and Lady Bacon
addresses him a letter, in which there are some allusions to Francis,
which will be read with interest:—
“This one chiefest counsel your Christian and natural mother doth give you even
before the Lord, that above all worldly respects you carry yourself ever at your first
coming as one that doth unfeignedly profess the true religion of Christ, and hath
the love of the truth now, by long continuance, fast settled in your heart, and that
with judgment, wisdom, and discretion; and are not afraid or ashamed to testify
the same by hearing and delighting in those religious exercises of the sincerer sort,
be they French or English. In hoc noli adhibere fratrem tuum ad consilium aut
exemplum....
“I trust you, with your servants, use prayer twice in a day, having been where
reformation is. Omit it not for any. It will be your best credit to serve the Lord duly
and reverently, and you will be observed at first now. Your brother is too negligent
herein, but do you well and zealously; it will be looked for of the best-learned sort,
and that is best.”